熱帶森林中具有豐富的生物多樣性，其中也包含兩棲動物。 然而全球的兩棲動物的族群正因棲地破碎或消失、全球氣候變遷、過度利用、和疾病等原因正急遽的下降, 因此測量其生物多樣性及鑑定稀有和常見物種在群聚生態學和保育上是相當必要的工作。 越南中部的原生兩棲動物的生物學及生態學目前大多都是未知的，而且其物種的組成,多樣性、和分布的資訊也是缺乏的。 其中多疣棘蛙 [Quasipaa verrucospinosa (Bourret, 1937); Granular Spiny Frog] 就是由於森林和溪流棲地的消失、環境劣化、和人類過度利用而頻臨滅絕。
本文的研究目的是 (1) 評估原始森林和次生林中兩棲動物的組成及各物種的數量，並藉此資料評估物種多樣性 (species diversity)：物種的豐度 (richness)、異質性 (heterogeneity)、和均勻度 (evenness)，並計算α、β、和γ多樣性。 此外，本研究也評估 (2) 多疣棘蛙的棲地佔據率和分布，並說明其族群生態和生活史特徵，包含食性組成、時空與體型有關的變異、繁殖生物學的季節模式、雄蛙鳴叫聲的特徵和變化、及蝌蚪變態的模式。
本研究的結果指出，兩區域兩棲動物物種的組成在地景尺度的相似度相當高 (0.86)，但在原始森林和次生林間則較低 (0.03)。 本研究共捕獲13,529隻的個體，分別屬於2目 (Anura and Gymnophina)、8科、33屬、和101種的物種。 次級森林中的兩棲類動物會包含一部分初級森林的特徵 (28.7%)。 原始森林中的物種豐富度和數量顯著高於次生林中 (P < 0.001)。 初級森林的兩棲類物種比次級森林豐富。 理論上，從整個生態系中，我們可以透過觀察指出整個生態系大約由106種組成，其中約有低於4.48物種數不屬於這個群集。
根據多次多疣棘蛙的棲息地數據檢測，從最好的樣本中我預估棲息率達0.632，比原始的棲息所得到的0.403多出57%。 初級森林的變因是造成棲息率變動很大的因素。 但是對於次級森林來說，其變因卻不會造成棲息率太大的改變。 生物檢測的模組[p(溫度、濕度、降雨量)]包含90.9%的總重，提供很明顯的證據指出環境因素對於模擬檢測機率是很重要的。
以出現頻度 (frequency of occurrence) 和體積百分比 (percent volume) 表示食性的結果顯示，多疣棘蛙共利用了2645項、27目、和9綱的物種，其中以無脊椎動物為主，另外也包含魚類、青蛙、及同種的亞成體。 無脊椎動物的種類有蜘蛛、甲蟲、螃蟹、膜翅目 (hymenopterans)、蚱蜢、蟋蟀、和蟬，其中單昆蟲就佔了59.8%的重要性。 平均每月消耗獵物體積和溫度成正比，和降雨量成反比。消耗獵物的體積和吻缸長 (snout-vent length) 及嘴寬度有正相關，此結果支持嘴基限制假說 (gape limitation hypothesis)。
多疣棘蛙的雄性在出生9個月後達到性成熟時，吻缸長為98.36 mm，雌性則在出生11月後達到性成熟，吻缸長為109.45 mm。 經組織學分析顯示，雄性的精子並非終年皆有，因此成年雄性的繁殖活動並非式連續性的；而繁殖的雌蛙也只在雨季 (9月 ~ 12月) 被觀察到，因此雌蛙的繁殖是有季節性的。 雌性的生殖線從七月開始增加體積，在雨季時有最多的產蛋量，其產卵數和其吻缸長有顯著關係，且在季節和不同月份間有變化。 經複迴歸 (Multiple regression analyses) 的分析顯示睪丸和卵巢的體積只受雨量顯著正向的影響，而不受溫度的影響。
多疣棘蛙的鳴叫聲可分為宣告型 (advertisement) 和侵略型 (aggressive) 兩種。 宣告型的主頻率 (dominant frequency) 為836 Hz，而侵略型的主頻率則在1020 Hz。 鳴叫持續時間 (call duration)、脈衝次數 (number of pulses)、脈衝率 (pulse rate)、脈衝間隔 (pulse interval) 在個體間有顯著的差異，然而主頻率是最重要的特性（一種統計特性)。 脈衝長度在個體間有中度差異。 雄性體型大小 (以吻缸長表示) 會顯著影響主頻率，此結果支持體型大小會限制鳴叫特徵的假說。
每次產卵平均產出有518隻蝌蚪，蝌蚪的存活率在變態的完成階段為80%，而變態過程需55.8天。 複迴歸分析結果顯示，水溫、溶氧量、及酸鹼值 (pH value) 會顯著影響蝌蚪的存活率和變態的天數。 在變態過程中的第46天，亞成蛙重2.3 g，吻缸長為25.8 mm。 Two-way ANOVA的分析結果顯示，蝌蚪體型大小在年間、地點、和年與地點的交互作用有顯著變化。

Tropical forests have rich biodiversity, including many amphibian species. However, amphibian declines are a global phenomenon that is occurring at an alarming speed. Habitat fragmentation or loss, global climate change, over-exploitation, and disease are the main causes that have led to these declines. Thus, measuring biodiversity and identifying the rarity and commonness of species play essential roles in both community ecology and conservation. However, the biology and ecology of most native amphibians in central Vietnam are largely unknown, and information on amphibian species composition, diversity, and distribution is lacking. This includes the granular spiny frog Quasipaa verrucospinosa (Bourret, 1937), which is already classified as “Near Threatened” due to loss of forest and stream habitats, environmental degradation, and human exploitation.
This dissertation investigates species composition and the number of frogs of each amphibian species in primary and secondary forests. Based on these data, I estimated species diversity (i.e., species richness, heterogeneity, and evenness) as well as rarefaction curves, and determined the lognormal distribution of amphibian species. I also estimated site occupancy rates and the distribution of granular spiny frogs, and described certain aspects of the population biology and life history of this species. These include the dietary composition, spatiotemporal and size-dependent variation, seasonal patterns of reproductive biology, acoustic features and variation of male calls, and patterns of tadpole metamorphosis.
The results of this study indicate that the overall similarity of species composition of amphibians between the two regions (0.86) within a landscape scale was considerably higher than that between primary and secondary forests (0.03). I caught 13,529 individuals of 101 species in 33 genera, eight families, two orders (Anura and Gymnophiona). The amphibian community in secondary forests contained a number of species characteristic of primary rain forest habitat (28.7%). The species richness and relative abundance were significantly higher in primary forests than in secondary forests (P < 0.001). The expected number of amphibian species in primary forests was significantly richer than in secondary forests. The expected total number of species theoretically available for observation in the entire community (including the rare species not yet collected) indicated that the “universe” comprises approximately 106 species, with the number of species predicted to fall below the veil line about 4.48 species.
Based on the detection/non-detection data for each site over multiple visits for granular spiny frogs, from the best model among all candidate models, I estimated a site occupancy rate of 0.632 that was higher than the naïve occupancy estimate of 0.403 and a 57% increase over the rate of sites at which frogs were actually observed. The site variable of primary forest was an important determinant of site occupancy, whereas occupancy was not associated with the variable of secondary forest. The species detection model p(temperature, humidity, rainfall) included 90.9% of the total weight, providing clear evidence that environmental conditions were important sample covariates in modeling detection probabilities.
Dietary analysis of granular spiny frogs identified 2645 prey items from 27 orders and nine classes. Prey of granular spiny frogs comprised mainly invertebrates, but also included fishes, frogs, and conspecific sub-adults. The major prey items as determined by frequency of occurrence, item count, and percent volume were spiders, beetles, crabs, hymenopterans, grasshoppers, crickets, and cicadas. Insects alone accounted for an importance value of 59.8%. The mean monthly prey volume consumed was positively and negatively correlated to temperature and rainfall, respectively. The volume of prey consumed was positively correlated with snout-vent length and mouth width of granular spiny frogs, supporting the gape limitation hypothesis.
Male frogs reached sexual maturity at 98.36 mm snout-vent length (SVL) and an age of nine months. Females reached sexual maturity at 109.45 mm SVL and an age of 11 months. Histological analyses revealed that sperm were not present throughout the year, indicating discontinuous reproductive activity for adult males. Reproductive females were observed only during months of the main rainy season (September to December), indicating that females reproduce seasonally. The gonads of females began to increase in volume in July and maximal egg production occurred during the main rainy season. Clutch size was correlated with female SVL and varied between seasons and months. Multiple regression analyses indicated that precipitation, but not temperature, was associated positively with testicular and ovarian volume.
Analysis of acoustic parameters revealed that granular spiny frogs emitted two call types (advertisement and aggressive calls). Advertisement calls had a mean dominant frequency of 836 Hz and aggressive calls had a mean dominant frequency of 1020 Hz. Call duration, number of pulses, pulse rate, and pulse interval were highly variable within individuals (dynamic properties), whereas the dominant frequency was the most stereotyped property (a static property). Pulse length exhibited an intermediate level of variation within individuals. Male SVL negatively influenced dominant frequency, supporting a body-size constraint for this call characteristic.
The average number of tadpoles per clutch was 518 individuals, the average survival rate at the complete stage of metamorphosis was 80%, and the total time to metamorphosis was 55.8 days. Multiple regression results for possible effects of water temperature, dissolved oxygen, and pH values on survival rates and the total time of metamorphosis of tadpoles were significant among localities. The process of metamorphosis is completed in stage 46, at which juvenile frogs have a mean body mass of 2.3 g and a mean SVL of 25.8 mm. Using a two-way multivariate analysis of variance revealed that body sizes of tadpoles varied significantly among years, sites, and by site-year interaction.

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